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Effects of the Blob on Settlement of Spotted Sand Bass, Paralabrax Maculatofasciatus, to Mission Bay, San Diego, CA

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Effects of the Blob on Settlement of Spotted Sand Bass, Paralabrax Maculatofasciatus, to Mission Bay, San Diego, CA RESEARCH ARTICLE Effects of the Blob on settlement of spotted sand bass, Paralabrax maculatofasciatus, to Mission Bay, San Diego, CA Anthony Basilio1, Steven Searcy1*, Andrew R. Thompson2 1 Environmental and Ocean Sciences, University of San Diego, San Diego, California, United States of America, 2 Fisheries Resources Division, Southwest Fisheries Science Center, NOAA Fisheries Service, La Jolla, California, United States of America * [email protected] a1111111111 a1111111111 a1111111111 a1111111111 Abstract a1111111111 The West Coast of the United States experienced variable and sometimes highly unusual oceanographic conditions between 2012 and 2015. In particular, a warm mass of surface water known as the Pacific Warm Anomaly (popularly as ªThe Blobº) impinged on southern California in 2014, and warm-water conditions remained during the 2015 El Niño. We exam- OPEN ACCESS ine how this oceanographic variability affected delivery and individual characteristics of lar- Citation: Basilio A, Searcy S, Thompson AR (2017) val spotted sand bass (Paralabrax maculatofasciatus) to an estuarine nursery habitat in Effects of the Blob on settlement of spotted sand southern California. To quantify P. maculatofasciatus settlement patterns, three larval col- bass, Paralabrax maculatofasciatus, to Mission Bay, San Diego, CA. PLoS ONE 12(11): e0188449. lectors were installed near the mouth of Mission Bay, San Diego CA, and retrieved weekly https://doi.org/10.1371/journal.pone.0188449 from June±October of 2012±2015. During `Blob` conditions in 2014 and 2015, lower settle- Editor: Heather M. Patterson, Department of ment rates of spotted sand bass were associated with higher sea surface temperature and Agriculture and Water Resources, AUSTRALIA lower wind speed, chlorophyll a (chl a) and upwelling. Overall, the number of settlers per Received: July 13, 2017 day peaked at intermediate chl a values across weeks. Individual characteristics of larvae that settled in 2014±2015 were consistent with a poor feeding environment. Although set- Accepted: November 7, 2017 tlers were longer in length in 2014±15, fish in these years had slower larval otolith growth, a Published: November 27, 2017 longer larval duration, and a trend towards lower condition, traits that are often associated Copyright: This is an open access article, free of all with lower survival and recruitment. This study suggests that future settlement and recruit- copyright, and may be freely reproduced, ment of P. maculatofasciatus and other fishes with similar life histories may be adversely distributed, transmitted, modified, built upon, or otherwise used by anyone for any lawful purpose. affected in southern California if ocean temperatures continue to rise in the face of climate The work is made available under the Creative change. Commons CC0 public domain dedication. Data Availability Statement: All relevant data are within the paper and its Supporting Information files. Introduction Funding: Funding was provided by a faculty research grant from the University of San Diego to A central goal of marine ecology and fisheries biology is to understand factors that contribute Steven Searcy as well as from support by the to variability in population size [1]. For many marine organisms understanding causes of fluc- Department of Environmental and Ocean Sciences tuations in abundance is complicated by a two-part life history with pelagic larvae that develop at the University of San Diego. The funders had no role in study design, data collection and analysis, offshore and settle to benthic juvenile/adult habitats [2±5]. The number of larvae that reach decision to publish, or preparation of the settlement is a critical factor contributing to future year-class strength [3, 6±8], although post- manuscript. settlement processes can also be an important source of variation [9, 10]. PLOS ONE | https://doi.org/10.1371/journal.pone.0188449 November 27, 2017 1 / 15 Settlement of spotted sand bass, Paralabrax maculatofasciatus Competing interests: The authors have declared Settlement success is dependent on a variety of processes including egg production [11, 12] that no competing interests exist. physical transport and retention of larvae [6, 13±18], and larval mortality [19]. Even small vari- ations in larval mortality can lead to large changes in the number of individuals that survive this period [20]. Larval mortality may be caused directly by predation [21], as well as indirectly through environmental conditions such as water temperature and food supply [22±27]. Food availability is particularly important to larval survival because larvae have high metabolic rates and low energy reserves [28]. At elevated water temperatures, metabolic rates are faster, thereby increasing energetic requirements and increasing risk of starvation [22, 23, 25, 29, 30]. After only a few days without food, the larvae of some fish species reach a "point of no return" and are unable to survive [31±33]. Water temperature and food availability may also affect larval survival by influencing growth rates and physiological condition [34±37]. The growth mortality hypothesis [38] sug- gests that if mortality is lower for larger individuals, then faster growing individuals of a given age will have a lower probability of mortality than slower growing individuals of the same age [39]. Larger individuals may be able to detect and respond to predators more effectively [21, 40], obtain food and withstand starvation [21, 41, 42], although, some studies suggest that larger larvae may actually have higher mortality [39, 43±45]. Finally, larvae with faster growth may be exposed to overall lower levels of predation by quickly growing out of this vulnerable life history stage [38, 46, 47]. Increased sea surface temperature (SST), may also be related to poor larval growth condi- tions by indicating periods with low upwelling and potential stratification of the water column [48, 49]. During upwelling events, increased supply of nutrients to the surface water can spur phytoplankton growth and support a trophic pyramid that includes zooplankton and many fish species [50]. When upwelling ceases, SST and stratification increase, and there is decreased productivity [48]. Indeed, previous work indicates that during a long-term shift from a cool to a warm water regime as the result of changing Pacific Decadal Oscillation there was an overall 46% reduction in volume of phyto- and zooplankton and a corresponding decline in abun- dance of many species of larval rockfish (Sebastes sp.) [51]. Understanding how settlement is affected by environmental conditions such as water tem- perature is especially important to gain insight into how warming ocean conditions may impact marine populations. In the northeast Pacific Ocean, during the winter of 2013±14, weakened winds and unusually high sea level pressure formed a region where heat was retained in surface water. This mass of warm water, known as ªthe Blobº, reached coastal waters in the western United States in the spring/summer of 2014 [52] and resulted in wide- spread changes in the biological structure and composition of both open-ocean and coastal ecosystems [53]. Sea surface temperatures were 1±4ÊC higher than average along the west coast of North America [53]. A geographically distinct section of the Blob near San Diego was named the Southern California Warm Anomaly (SCWA) and increased thermal stratification which led to a reduction of vertical mixing and nutrient fluxes into the surface water [54]. This pool of warm water was partially responsible for the shift in the California Current from a pro- ductive La Niña state in 2013 to a warm area with low productivity [55]. Our objective was to examine how changes in environmental conditions over four years (2012±2015) affected daily and annual settlement success of spotted sand bass (Paralabrax maculatofasciatus) in Mission Bay, CA. We also examined how environmental conditions influenced individual characteristics at settlement (length, age, condition, larval growth rates). The sampling years included two years of relatively normal conditions (2012±13) followed by the arrival of the SCWA in 2014 and dramatic changes in chlorophyll a (chl a) and SST that continued through 2015. PLOS ONE | https://doi.org/10.1371/journal.pone.0188449 November 27, 2017 2 / 15 Settlement of spotted sand bass, Paralabrax maculatofasciatus Materials and methods Ethics statement Sampling was conducted under California Department of Fish and Wildlife scientific sampling permit SC-11846 with the approval of the University of San Diego's Institutional Animal Care and Use Committee (IACUC). Study species Spotted sand bass (Paralabrax maculatofasciatus) are a recreationally important fish in South- ern California typically found from Santa Monica Bay, CA in the north to Mazatlan, Mexico in the south [56]. There are two main populations of P. maculatofasciatus along this range: one in the Gulf of California and another in the Pacific along Baja California and Southern California [57]. There is likely a high degree of larval retention as P. maculatofasciatus sampled near San Diego are genetically distinct from those further south along the Baja California peninsula [57]. Spotted sand bass adults are primarily found in calm, shallow, nearshore habitats such as bays and estuaries [56]. Adults spawn near the entrance of bays in the late spring and summer [56], are capable of spawning multiple times throughout the season and may even spawn daily [58]. Larvae develop in the
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